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Seed collection and germination strategies for common wetland and coastal sage scrub species in Southern California.

Abstract.--There is a need for a consolidated source of information on native vegetation seed collection and germination strategies in southern California. Published literature on these methods is often experimental, species-specific, and widely scattered throughout online and print media. Planting and restoration strategies may need to be site-specific; however, similar methodological approaches are often utilized allowing for the development of general strategies for seed collection, storage, and germination methods. A better understanding of species-specific seed attributes and growth processes will help restoration ecologists collect high-quality, viable seed, thereby increasing the potential success of the restored vegetation community by reducing plant mortality, project costs, and effort. This paper synthesizes seed collection and germination strategies for native vegetation common to southern California estuarine wetland, coastal dune, and coastal sage scrub systems.

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Current estimates affirm that over 70% of coastal wetlands in the Southern California Bight have been lost since the 1800's, with estimates increasing to over 95% for highly urbanized areas, such as Los Angeles County (Stein et al. 2014). The magnitude of these losses and the continued degradation of coastal wetland systems, and adjacent upland and coastal sage scrub habitats, threatens the ecological integrity and sustainability of these habitat types and their watersheds. To address these issues, a number of restoration and mitigation projects aimed at restoring lost ecosystem services, increasing biodiversity, boosting resilience, and in the case of mitigation, creating new wetland habitat, are currently in the planning process in southern California (Noss 2000, Zedler 2000). The majority of wetland restoration or mitigation projects develop a site-specific framework of protocols and management strategies outlining a planting and re-vegetation strategy.

Planting strategies designed to establish self-sustaining plant communities identify both the species to be included in the restoration and the source of plant material (i.e. nursery stock or local seeds) (Zedler 2001). Restoration plant palettes should be designed to mimic reference or historic site diversity and be composed of an appropriately broad range of species (Zedler 2001, Johnston et al. 2012). Because of their unique location in the landscape as the connecting habitat between marine, terrestrial, and freshwater ecosystems, coastal wetland complexes naturally support a variety of salt marsh, brackish, and freshwater plant species (Lichvar et al. 2014). Species from each of these habitat types should be incorporated into an appropriate plant palette. Evidence also suggests that the wetland-upland ecotone should be considered an extension of wetland habitat for conservation and restoration purposes (James and Zedler 2000, Wasson and Woolfolk 2011). Thus, coastal sage scrub, dune, and transitional species commonly found in the wetland-upland ecotone should be considered in wetland restoration re-vegetation strategies.

Once a restoration plant palette has been developed, species-specific plant material (e.g. seeds and seedlings) acquisition and propagation methods must be determined. While plant material can be obtained from local nurseries, collection and propagation of native seed from local sites is considered the most cost-effective and ecologically-sound method of sourcing germplasm for restoration and mitigation projects (Zedler 2001, Broadhurst et al. 2008). Site-specific or nearest neighbor collections are preferred to distant collections and use of nursery stock, as locally-collected individuals are better adapted to community environmental conditions, maintain local genetic integrity, ensure persistence of local eco-types, prevent unintended gene flow, may improve the long-term sustainability of the site, and may enrich the diversity of the wetland plant community (Guerrant 1996, Montalvo et al. 1997, Bowler 2000, Zedler 2001, Mitsch and Gosselink 2010, Vander Mijnsbrugge et al. 2010). Non-local genotypes may be maladapted to local site conditions, leading to improper establishment, or negative impacts to plant and animal communities through competition or species hybridization (Bischoff et al. 2006, Vander Mijnsbrugge et al. 2010). The retention of local eco-types and genetic information is gaining importance in the field of restoration biology, reflected by the recent inclusion of required onsite and/or near neighbor collections by regulatory agencies overseeing restoration and mitigation work (Bowler 2000). It is also important to note that making collections in many nature preserves requires permits and/or express permission from the regulating agency.

A number of techniques exist to propagate plant material for wetland, coastal sage scrub, and dune species. Seeds are often the primary means of reintroducing native plant species to restoration sites in a number of habitat types (Montalvo et al. 2002, Merritt and Dixon 2011). Restoration sites may be seeded using a variety of techniques (e.g. broadcast seeding, drilling, imprinting, or hydroseeding) or collected, cultivated in a greenhouse, and transplanted to the site (Bowler 2000, Montalvo et al. 2002, Merritt and Dixon 2011). Simple seeding experiments generally are performed with limited success, especially at lower elevations or within tidal wetland habitats, as seeds often fail to germinate or float away with rising tides (Broome et al. 1988, Zedler 2001). Techniques like hydroseeding that involve mixing seed with water and either mulch, soil, or organic matter prior to application, tend to work well for many wetland and coastal sage scrub species [e.g. Salvia mellifera (black sage) and Eriogonum fasciculatum (California buckwheat)] and may enhance seedling establishment (Zedler 2001, Montalvo et al. 2002, Montalvo and Beyers 2010).

Transplanting greenhouse-grown seedlings is an effective re-vegetation strategy that may increase the potential establishment success when compared to direct seeding for some species. In one experiment, survivorship of 2-4 month old marsh seedling transplants was over 95% for all but one treatment, much higher than the success rate of direct seeding (Zedler 2001). Seedlings of a variety of halophytic marsh species including Suaeda esteroa, estuary seablite, and Salicornia bigelovii, dwarf pickleweed, and a variety of coastal sage scrub species like Atriplex canescens, four-wing salt bush, have been successfully grown in greenhouses and transplanted for restoration purposes (Zedler 2001, Francis 2009). While use of seeds and seedlings has been successful for many species [e.g. Achillea millefolium (common yarrow) and Astragalus tener var titi (coastal dunes milk vetch)], effective propagation techniques are species-specific and other species, like Batis maritime, saltwort, do not readily grow from seed and require use of alternate methods (Zedler 2001).

Other common approaches to generate plant stock include use of cuttings, root division, and direct transplantation of seedlings or mature plants to the site of interest (Zedler 2001, Baskin and Baskin 2014). Direct transplantation of coastal sage scrub seedlings [e.g. Artemisia californica (California sagebrush), Salvia mellifera, Encelia californica (California brittlebush), and Eriogonum fasciculatum] and mature plants salvaged from donor sites have been used with great success in mitigation efforts (Bowler et al. 1994, Bowler 2000). Similarly, use of transplants, sod, and small plugs of wetland soil, have been effective in introducing a number of wetland species, including Spartina foliosa, California cordgrass, to sites (Tmka 1998, Zedler 2001, Mitsch and Gosselink 2010). Use of cuttings is documented to work well for other species; cuttings of Salicornia pacifica, common pickleweed, for example, have been successfully propagated by Tree of Life Nursery in San Juan Capistrano, California. While each of these approaches has merit, the discussion in the remainder of this paper (and the accompanying appendices) focuses on the use of seeds and greenhouse-grown seedlings to target a data gap in peer-reviewed literature.

While general techniques for successfully establishing common wetland and coastal sage scrub species described in the preceding paragraphs are understood (Broome et al. 1988), the field of restoration biology is still developing and could benefit greatly from additional research. More specifically, the field could benefit from research regarding species-specific collection and propagation techniques because cultivation and planting strategies are often species-specific, highly variable, proprietary, or experimental. Information for many native species of interest does not exist, or is not publically available, forcing restoration managers and ecologists to rely on general information about the genus or costly and time-intensive exploratory studies (Dreesen and Harrington 1997). Publically available sources are scattered throughout a variety of peer-reviewed and non-peer-reviewed resources. With over a dozen wetlands in southern California considered candidates for large-scale wetland restoration projects, a compilation of literature summarizing re-vegetation strategies for the region is needed (SCCWRP 2001). This paper synthesizes basic seed characteristics, as well as collection and germination strategies for vegetation species common to estuarine wetland and adjacent upland habitat types, specifically coastal salt marsh and coastal sage scrub habitats in southern California.

Materials and Methods

Common seed collection, germination, and propagation techniques are described in the text of this paper. General species information (e g. scientific name, common name, and habitat type) is included in Appendix I. Detailed species-specific data and recommendations are included in Appendix II, which summarizes available information for 66 native plant species commonly used in southern California coastal restoration projects. Species-specific details were compiled using available literature. While the majority is derived from peer-reviewed publications, some non-peer reviewed literature was included to fill data gaps in published information. As many data gaps exist, and gray literature was used throughout the article text and the accompanying appendices, the authors have chosen not to distinguish gray literature with footnotes and this was approved by the editors. Instead, these sources are listed, with all peer-reviewed sources, in the Literature Cited section. In instances where duplicate information was identified, the source with the most extensive experimental results was cited. Field observations from the Ballona Wetland Ecological Reserve, Los Angeles, CA, were used to determine some seed collection windows. Appendix II is not intended to be comprehensive; instead, it focuses on common coastal wetland and upland species in southern California for which there was available literature. Priority was given to information specific to southern California coastal habitats, but species-specific information from other geographic areas was included as needed for completeness. Implementation of specific methods may vary slightly by site or project. A number of resources exist that provide general species profiles of the plants described in Appendices 1 and II. Three websites in particular, S&S Seeds (http://www.ssseeds.com), the Theodore Payne Foundation (http://theodorepayne.org), and Tree of Life Nursery (http://www.califomianativeplants.com), are recommended for supplemental information relating to life history and planting recommendations.

Materials

Equipment and supplies needed for seed collection, cleaning, and germination are highly variable based on the specific vegetation species. Recommended field, laboratory, and greenhouse equipment are listed in Table 1. In addition to the field equipment listed, available background information (e.g. reports, vegetation maps, taxonomic keys) should be brought into the field to aid correct taxonomic identification of species.

Seed Collection

Seeds should be collected within seed zones, geographic zones in which genetic exchange naturally occurs. Practitioners are advised to use life history traits, landscape context, and available genetic studies to correctly determine seed zones (Krauss and He 2006). It is important to note that due to extensive urbanization and fragmentation in southern California, historic areas of seed exchange have been diminished. In addition to considering provenance of seeds, care should be taken to ensure that seed collections contain sufficient genetic diversity (Vander Mijnsbrugge et al. 2010) as diversity safeguards against disease, environmental fluctuations, and inbreeding depressions (Smith et al. 2007). To maximize the range of genetic diversity represented in the collection, seed should be collected from 10-50 individuals per population (Lippitt et al. 1994, Vander Mijnsbmgge et al. 2010). Local adaptations and site-specific variability should also be taken into consideration, but site-specific recommendations are outside the scope of this product. When collecting seeds, less intense and more frequent seed harvests are preferable to infrequent and intense harvests (Wall 2009). Negative impacts on the seed source population must be considered (Krauss and He 2006). A general safe harvesting recommendation is to take no more than 5% of seed from a given species and geographic area (Zedler 2001).

Once plant identity has been confirmed, carefully examine the seeds to assess maturity. Avoid collection of immature seed, as premature collection may result in low seed viability (Bonner and Karrfalt 2008, Baskin and Baskin 2014). In general, it is good practice to begin collecting seeds around the time that natural dispersal begins (Baskin and Baskin 2014). Seeds are considered ripe if seed capsules are dry and tan or brown in color, rather than yellow or green (Lippitt et al. 1994, Bonner and Karrfalt 2008, Baskin and Baskin 2014). Frequent visits to collection sites are suggested to repeatedly assess seed stage within the recommended collection time window. For species with insufficient published seed collection data or information, e.g. Artemisia douglasiana, detailed field notes are essential to pinpoint the ideal collection window and successfully collect seeds.

Once the seeds of target species are deemed ripe, the collection process can begin. Collection/ isolation of seed varies based on plant anatomy. Observe the plant and note if the species has berries or dry fruits, dehiscent or indehiscent seeds, and note if seeds are in seed heads or seed clusters as collection methods vary for each category (Table 2). Additionally, if a species is known to be dioecious [e.g. Croton californicus (California croton), Baccharis spp., Salix spp.], care should be taken to ensure that sufficient seed quantities are collected from both male and female plants (Clarke et al. 2007). Vouchering specimens from collected seeds is a good practice and should be considered during the planning phase.

Seed Cleaning

Seed cleaning removes floral parts, seed coats, pods, fleshy fruit material, and other debris from seeds (Jorgensen and Stevens 2004). Machinery, including aspirators, hammermills, fanning mills, and blowers, exists to aid large-scale seed enterprises. Hammermills, fanning mills, and blowers help isolate seed and remove chaff and floral parts (Shaw 1975, Jorgensen and Stevens 2004). Although seed cleaning machinery is useful, cleaning for small-scale projects can be efficiently performed by hand (Bonner and Karrfalt 2008). To isolate seeds and remove excess chaff, remove seeds from branches and large floral parts. Then, rub remaining seeds and floral parts over a sieve. Once seeds are isolated from chaff, only retain seeds that look healthy and ripe (i.e. dark brown/tan in color, fully-formed). For some species, chaff does not present a huge problem, and it may be more efficient to seed with some chaff. Discard seeds that appear sickly or deformed. If the seed is contained in a capsule, gently crush the capsule by hand or with a rolling pin. Removal of woody capsules, as seen in Abronia spp., may also be aided with the use of generic nail clippers (RM. Drennan, personal communication).

Seed Storage

For the greatest germination yield, storage time should be minimized, and use of newer seeds should be prioritized. While native seed longevity varies by genus and species, a number of seeds are known to be short-lived. For example, seeds of Lycium californicum, California box-thorn, are viable for up to one year at most. While seeds of other species [e.g. Atriplex spp., Astragalus spp., and Lupinus chamissonis (dune bush lupine)] will remain viable for much longer (i.e. 4-10 years), the germination rate of seeds in long-term storage will likely decline over time. In addition to reducing germination rate, long-term storage will often induce seed coat or embryo dormancy, and stored seeds may need to be treated prior to planting. For example, the hard seed coat of Astragalus tener var. titi seeds may require scarification, or mechanical scraping with sandpaper, a file, or a knife, to initiate germination if stored for an extended period of time (Baskin and Baskin 2014, USFWS n.d.)

The longevity of certain seeds can be increased if best management practices for storage are followed for the species and/or general seed storage procedures are applied. Most dry seeds should be stored at low temperatures, 10-15.6[degrees]C (50-60[degrees]F), and low humidity, less than 40% relative humidity (Jorgensen and Stevens 2004, Recon Native Plants Inc. 2015). Substandard storage in conditions with fluctuating temperatures or high humidity may result in significant seed loss (Merritt and Dixon 2011).

Germination Considerations

Successful propagation of southern California coastal plant species requires a thorough understanding of seed germination ecology. Seed germination is dependent upon a number of evolutionary and ecological factors which generally must be observed, and often replicated, in the laboratory or greenhouse to successfully grow propagules. These, often species-specific, factors include, but are not limited to: germination timing/seasonality, environmental conditions, such as temperature, soil texture, soil moisture, soil salinity, light availability, presence of smoke, and seed age, and dormancy state, both at the time of maturation and dispersal (Baskin and Baskin 2014).

Germination Timing

Seeds are adapted to germinate under favorable environmental conditions (Deberry and Perry 2000). An understanding of natural germination timing is helpful in determining the environmental conditions that best promote germination of a particular species in the greenhouse or laboratory. This is particularly true, as in both the greenhouse and laboratory, environmental conditions can be manipulated to mimic natural seasonal variation. Temperature, moisture, and light are generally controlled for this purpose (see 'Temperature' and 'Light' sections below) (Noe and Zedler 2001).

Temperature

Understanding germination timing under natural conditions will often indicate what range of temperatures best promote germination. Temperature influences germination directly through regulation of enzymatic reactions, or indirectly by controlling the synthesis of hormones that alter seed dormancy. While temperature is an important determinant in the regulation of both germination and dormancy, response to temperature in freshwater wetland species seems to be dependent on habitat, not phylogenetic relatedness. Temperature interplays with other environmental conditions to promote germination (Brandel 2006). Further, the germination rate of certain species is enhanced with simulated temperature fluctuations, rather than constant temperatures. While response to fluctuating temperatures depends both on specific species and habitat, a few generalities exist. Both small-seeded species and forbs tend to respond well to fluctuating temperatures while larger-seeded and graminoid species do not show as marked a preference for temperature fluctuations (Liu et al. 2013).

Soil Texture

To grow seedlings, clean, viable seeds should be planted in mixtures of sand, top soil, and peat moss or vermiculite (Broome et al. 1988). To achieve the greatest germination rate, the exact composition of the mixture should be tailored to the individual plant species of interest. Life history and preferred habitat of the species should be considered when determining optimal soil conditions. For instance, Abronia maritima, which naturally occurs on sandy dunes, should be sown in soil consisting largely of sand, or other coarse grains.

Soil Moisture

Soil moisture must also be considered when sowing seeds (Noe and Zedler 2000, Noe and Zedler 2001). Most mature seeds must imbibe in the early stages of germination to activate enzymes (Deberry and Perry 2000). After seeds imbibe, sufficient, and relatively constant soil moisture is needed to ensure proper germination (Bonner and Karrfalt 2008). Most species in southern California salt marsh systems germinate well in moist soil at low salinity (Zedler 2001). Experiments suggest that Distichlis spicata grows best with a fluctuating inundation regime, where inundation was varied over time, but the soil surface was never completely dry (Elsey-Quirk et al. 2009). Germination of other high marsh plant species is highest with 41-51% soil moisture (Zedler 2001).

It is important to note that while seeds of wetland species are adapted to wet conditions with limited oxygen, coastal sage scrub and upland transition species are more sensitive to inundation. For these species, excessive exposure to water can be problematic, causing seeds to become waterlogged (Fenner 1992, Deberry and Perry 2000). Following germination, water regimes, that specify both the quantity and frequency of water application, both in the greenhouse and in natural environments, may influence growth rates and should be carefully considered.

Soil Salinity

Another major factor that influences germination is soil salinity (Noe and Zedler 2000, Noe and Zedler 2001). Certain halophytic species, like Salicornia bigelovii, germinate to higher percentages under somewhat saline conditions (0.05-0.09 M). In general, although halophytes are salt-tolerant, high percentages of halophyte seeds will germinate in distilled water. Results of salinity experiments suggest that seeds will often germinate to higher percentages in distilled water, as seeds tend to be sensitive to salt concentrations, and exposure to excessive salt can drastically decrease germination yields. Still, much variation exists in the germination of halophyte species in saline environments (Baskin and Baskin 2014).

Light

Light is another environmental factor that affects germination. Exposure to light is often required for germination to occur. Exposure to light has been documented to improve germination rates for certain species [e.g. Eriogonum fasciculatum (California buckwheat), Baccharis salicifolia (mule fat)] (Zedler 2001, Bonner and Karrfalt 2008). Still, exposure is not always sufficient to ensure the successful occurrence of germination mechanisms. Duration of exposure to light (i.e. day length or photoperiod) also plays an important role in seedling emergence and growth of southern California natives (Sprague 1944, Noe and Zedler 2000, Greiner and Kohl 2014). For instance, long-day conditions (16 hours of light for every 8 hours of darkness) are necessary to successfully culture Oenothera species (Greiner and Kohl 2014). Photoperiod may also influence other processes, such as flowering. Melica imperfecta and Stipa lepida have been shown to flower 10-20 weeks faster with constant light (i.e. 24-hour photoperiod) when compared to an 8-hour photoperiod (Ashby and Flellmers 1959).

Smoke Treatments

Southern California, like most regions with Mediterranean climates, is subject to frequent and intense wildfires, and certain species have adapted to be fire-tolerant (Keeley and Fotheringham 1998, Crosti et al. 2006). Germination of fire-tolerant species is generally enhanced by exposure to fire or smoke (Crosti et al. 2006, Baskin and Baskin 2014). Smoke-stimulated germination, via exposure to liquid or aerosol components of smoke, may be useful for many coastal sage scrub species. For instances, exposure of Salvia mellifera seeds to smoke or other components of fire, like charred wood or potassium nitrate (KN[O.sub.3]), may help stimulate germination (Montalvo and Beyers 2010).

Other Considerations

In some instances, information regarding the necessary conditions or procedures to promote germination is not readily available for a particular species. In such situations, it is advisable to consult local experts that may have species-specific knowledge. Alternatively, simple tests or experiments manipulating a variety of the environmental factors discussed above may be performed.

Germination Testing

If a seed lot requires germination studies, it is preferable that they are conducted shortly after seed collection, within 7-10 days, to ensure seeds are viable and have not entered seed dormancy. Germination trials can test outcomes of various pre-treatments and/or growing conditions. They are often also used to express the quality of a seed lot (Lippitt et al. 1994). The results of germination trials are typically reported as percentage germination or germination rates. Percentage gennination is the percentage of seeds that germinate under the specified set of conditions. Comparing germination rates of a variety of treatments allows easy determination of the most effective combination of germination conditions.

While germination rates are useful, the industry will often use other terms to describe the percentage of seed that will germinate under a given set of conditions. Pure Live Seed (PLS) is a common way to express viability. PLS is calculated by multiplying the percentage of pure seed by the percentage of total viable seed and dividing the product by one hundred (S&S n.d., Showers 2010). Other measures include specification by purity, bulk pounds, or PLS pounds (S&S).

Dormancy Considerations

Seeds for a number of wetland plants are known to be dormant. In these species, seed dormancy must be broken to promote growth and germination (Baskin and Baskin 2014). The process is generally moisture and temperature dependent, but varies both with species and type of dormancy. Three types of dormancy should be considered: physical (or seed coat) donnancy, internal dormancy, and morphophysiological dormancy. Seeds with physical dormancy have seed coats or other structures that are impermeable to water and/or oxygen (Lippitt et al. 1994, Baskin and Baskin 2014). This form of dormancy is generally broken by penetrating/ opening the seed coat or specialized structure that excludes water or oxygen. This can be achieved through scarification, cold and warm stratification, or exposure to dry heat, charate, fire, acid, and light. Internal dormancy, caused by a physiological mechanism that inhibits germination, is generally broken through use of warm and/or cold stratification. Morphophysiological dormancy is similar to physiological, but seeds with this type of dormancy also have an underdeveloped embryo. A variety of methods can be used to break morphophysiological dormancy, including: scarification, submersion in hot water [82-93[degrees]C (180-200[degrees]F)], treatment with dry heat, exposure to fire, acid, mulch treatment, cold stratification, warm stratification, and exposure to light (Emery 1988, McClure 1997, Baskin and Baskin 2014). Common dormancy breaking methods are detailed in Table 3).

Unfortunately, as indicated by the variety of conditions listed above, there is not one prevailing standardized method to break seed dormancy. Again, methods vary based on the life history of the species. Species-specific life histories, available at the growers' websites listed above, can be a good indicator of the required conditions for that species. For example, species that typically germinate in early spring after a cold and/or rainy winter, such as Platanus racemosa, western sycamore, often require cold, moist stratification mimicking natural wintering to break dormancy. Other species, such as Acmispon glaber, common deerweed, require heat treatment to break dormancy which also correspond with the life history of that species; A. glaber does particularly well after wildfire events. However, treating seeds to break dormancy is not enough to guarantee germination. Germination requirements must also be considered. Methods and information should be supplemented by experimentation when necessary.

Mycorrhizae

Establishing functional ecosystems also requires consideration of subsurface components of the system. Many plants have symbiotic relationships with soil-inhabiting microorganisms, yielding root systems that are more effective at extracting water and nutrients from the rhizosphere (i.e. soil profiles influenced by root secretions and soil fauna). The fungus-root system is called mycorrhizae (Gerdemann 1968, Tree of Life Nursery n.d.). Research has shown that mycorrhizae can increase plant growth and are essential in successfully establishing vegetation during restoration and mitigation projects (Reeves et al. 1979, Allen and Allen 1980, Cooke and Lefor 1990). If planting areas are severely disturbed and lack a healthy rhizosphere, steps should be taken to ensure presence of mycorrhizae, or to increase the potential for natural development. As the presence of mycorrhizae is important in establishing many wetland and coastal sage scrub species, container plants are often inoculated prior to planting (Cooke and Lefor 1990, Bowler 2000). Seedlings can be inoculated with a spore suspension or via introduction of small amounts of collected soil from sites with a healthy rhizosphere to a sterile soil (van de Voorde et al. 2012). Starter-cultures are also available commercially.

Discussion

Southern California has lost a significant portion of its coastal ecosystems due to urban development, agriculture, invasive species, and in the case of coastal estuarine wetlands, severely modified hydrology resulting from both channelization and deposition of fill sediments (Westman 1981). Loss of these ecosystems is concerning because they provide valuable ecosystem services including supporting important fisheries, filtering water, sequestering carbon, and providing habitat for a diversity of plant and animal life, including a number of threatened and endangered species. Wetlands are buffered by transition habitats, and many wetland-associated species also require adjacent upland habitat areas to breed, roost, or to have the highest likelihood of survival. Plant species in Southern California also display a high degree of endemism and the Southern California coast is considered a global biodiversity hotspot (SCCWRP 2001).

Although wetlands in southern California have attained protected status and efforts are being made to restore degraded habitats throughout the Southern California Bight, the increasing human populations along the California coast will continue to impact these coastal ecosystems (Callaway and Zedler 2004). To preserve the spectrum of ecosystem services coastal wetlands and their adjacent upland habitats provide, managers throughout the southern California region need to work collectively to conserve remaining high quality coastal wetland habitat and to restore lower quality, degraded habitats.

Clearly, there is a significant and ongoing regional need for restoration projects to recover lost habitats and preserve the unique communities. Increased reliance on ecological restoration of vegetation assemblages emphasizes the need for sound, scientifically-tested techniques to ensure the successful reestablishment of plant communities. While this document is not comprehensive, and there is still a practical need for land managers to compile detailed site information and evaluate site-specific experiments prior to implementing a restoration scheme, this literature review compiles available seed collection and germination information for the southern California region and provides an initial assessment of published methods for common wetland, dune, and coastal scrub plants. Many unknowns remain in restoration ecology theory, and understanding of the most effective restoration practices remains incomplete. Knowledge gaps regarding the collection and germination requirements of integral species [e.g. Hazardia squarrosa (saw-toothed goldenbush)] and other species with limited research available [e.g. Elymus triticoides (creeping wild rye)] precluded their inclusion in this review. Planners are encouraged to conduct regular site monitoring and employ adaptive management strategies. In this way, progress can be evaluated and unexpected outcomes and shortcomings can be corrected.

Still, there is a regional need for additional research regarding seed phenology and maturation of southern California species. Although a number of wetland, dune, and coastal sage scrub restorations are planned in southern California, information regarding seed collection and germination for many naturally occurring species is not readily available. Therefore, the field of plant community restoration could benefit greatly from additional research regarding seed phenology and maturation, both in the form of species-specific experimentation and literature and broader-scale, regional or ecosystem-based reviews. Filling in existing knowledge gaps and developing a better understanding of seed processes will help restoration ecologists collect high quality, viable seed, thereby increasing the potential success of the restored vegetation community by reducing seed/seedling mortality, restoration cost and human effort.

Perhaps more importantly, the region could benefit from the development of a coordinated network of restoration ecologists. Compilation of this literature review suggests that information regarding the restoration of wetland plant communities is abundant, but it is dispersed, produced by various sources, and often proprietary. Intentional withholding of information by nurseries or private environmental consulting firms inevitably leads to duplication of efforts by groups working in the southern California region and surely impacts both the overall quality of restored habitats and project efficiency. Engagement and cooperation of existing private industry groups and public sector regulators with a vested interest in restoring coastal wetland plant communities would be a major victory and a tangible step forward for the threatened coastal ecosystems in the region.

While establishing vegetation in restored wetlands is a vital component to the overall restoration scheme, it is just a small part of the overall restoration process. Restoring wetland ecosystems is complex; plans must incorporate vegetation, hydrology, substrate, and marine and terrestrial animals. To fulfill restoration aims, well-informed, inter-disciplinary approaches that incorporate ecologists, engineers, managers, lawyers, and practitioners from other technical fields are needed (Zedler 2000, Kiehl 2010). Inter-disciplinary approaches will best foster creativity and progress knowledge and understanding in the field of restoration.

Acknowledgements

The authors wish to thank the colleagues and interns who assisted with this work. Special thanks to Phillippa M. Drennan, Remy G. Landon, and Jessica Sharpe whose contributions were significant. Thanks as well to Recon Native Plants, Inc. This work was made possible with financial support from the California State Coastal Conservancy and the US Environmental Protection Agency, but does not necessarily reflect their views and policies.

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Appendices
Appendix I. Species-specific habitat associations for wetland,
coastal sage scrub, and upland transition species common in southern
California. This table includes scientific and common names from
Jepson eFlora (http://ucjeps.berkeley.edu/, accessed June 4, 2015).
Habitat association information is derived from Jepson and further
refined with information available from the Manual of California
Vegetation (2nd edition), the S & S Seeds Plant Database
(www.ssseeds.com/plant-database), and the species-specific literature
cited in Appendix II (Baldwin et al. 2012, Sawyer et al. 2009, S&S
Seeds, n.d.).

Scientific name                      Common name

Abronia maritima                     Red sand verbena
Abronia umbel lata                   Pink sand verbena
Achillea millefolium                 Common yarrow
Acmispon glaber                      Deer vetch
Ambrosia psilostachya                Western ragweed
Artemisia californica                California sagebrush
Artemisia douglasiana                Mugwort
Artemisia tridentata                 Big sagebrush
Arthrocnemum subterminale            Parish's pickleweed
Astragalus pycnostachyus var.        Ventura marsh milk vetch
  lanosissimus
Astragalus tener var. titi           Coastal dunes milk vetch
Atriplex californica                 California orach
A trip lex canescens                 Fourwing saltbush
Atriplex lentiformis                 Big saltbush
Atriplex prostrata                   Fat-hen
Atriplex watsonii                    Watson's saltbush
Baccharis pilularis                  Coyote brush
Baccharis salicifolia subs,          Mule fat
  salicifolia
Baccharis sarothroides               Broom baccharis
Batis maritima                       Saltwort
Cressa truxillensis                  Alkali weed
Croton califomicus                   California croton
Distichlis littoralis                Shore grass
Distichlis spicata                   Salt grass
Encelia californica                  California brittlebush
Eriogonum fasciculatum               California buckwheat
Frankenia salina                     Alkali heath
Grindelia camporum                   Valley gum weed
Hazardia squarrosa                   Saw-toothed goldenbush
Heteromeles arbutifolia              Toyon
Hordeum brachyantherum               Meadow barley
Isocoma menziesii                    Coastal goldenbush
Iva axillaris                        Poverty weed
Jaumea camosa                        Fleshy jaumea
Juncus acutus subs, leopoldii        Southwestern spiny rush
Juncus bufonius                      Toad rush
Limonium californicum                Western marsh-rosemary
Lupinus chamissonis                  Dune bush lupine
Lycium californicum                  California box-thorn
Melica imperfecta                    Little California melica
Mimulus aurantiacus                  Sticky monkey
Oenothera elata                      Hookers evening primrose
Peritoma arborea                     Bladderpod
Phacelia ramosissima                 Branching phacelia
Plantago erect a                     Foothill plantain
Platanus racemosa                    Western sycamore
Populus fremontii subs, fremontii    Fremont cottonwood
Potentilla anserina subs, pacifwa    Pacific silverweed
Pseudognaphalium californicum        California cudweed
Rosa californica                     California wild rose
Salicomia bigelovii                  Dwarf pickleweed
Salicomia pacifwa                    Common pickleweed
Salix exigua subs, exigua            Narrow-leaved willow
Salix lasiolepis                     Arroyo willow
Salvia apiana                        White sage
Salvia mellifera                     Black sage
Schoenoplectus acutus var.           Common tule
  occidentalis
Schoenoplectus califomicus           Southern bulrush
Spartina foliosa                     California Cord grass
Stipa cemua                          Nodding needle grass
Stipa lepida                         Foothill needle grass
Suaeda esteroa                       Estuary seablite
Suaeda nigra                         Bush seepweed
Suaeda taxifolia                     Woolly seablite
Triglochin maritima                  Common arrow-grass
Vulpia microstachys var.             Small fescue
  microstachys

                                      Low     Mid    High    Salt
Scientific name                      marsh   marsh   marsh   pan

Abronia maritima
Abronia umbel lata
Achillea millefolium
Acmispon glaber
Ambrosia psilostachya
Artemisia californica
Artemisia douglasiana
Artemisia tridentata
Arthrocnemum subterminale              x       x
Astragalus pycnostachyus var.                  x       x      x
  lanosissimus
Astragalus tener var. titi                             x      x
Atriplex californica                                   x
A trip lex canescens
Atriplex lentiformis
Atriplex prostrata                                     x
Atriplex watsonii                                      x
Baccharis pilularis
Baccharis salicifolia subs,
  salicifolia
Baccharis sarothroides
Batis maritima                                 x       x      x
Cressa truxillensis
Croton califomicus
Distichlis littoralis                                  x      x
Distichlis spicata                             x       x
Encelia californica
Eriogonum fasciculatum
Frankenia salina                               x       x      x
Grindelia camporum
Hazardia squarrosa
Heteromeles arbutifolia
Hordeum brachyantherum
Isocoma menziesii
Iva axillaris
Jaumea camosa                          x       x
Juncus acutus subs, leopoldii
Juncus bufonius                                        x      x
Limonium californicum                          x       x
Lupinus chamissonis
Lycium californicum
Melica imperfecta
Mimulus aurantiacus
Oenothera elata
Peritoma arborea
Phacelia ramosissima
Plantago erect a
Platanus racemosa
Populus fremontii subs, fremontii
Potentilla anserina subs, pacifwa              x       x
Pseudognaphalium californicum
Rosa californica
Salicomia bigelovii                                    x      x
Salicomia pacifwa                      x       x
Salix exigua subs, exigua
Salix lasiolepis
Salvia apiana
Salvia mellifera
Schoenoplectus acutus var.
  occidentalis
Schoenoplectus califomicus
Spartina foliosa                       x
Stipa cemua
Stipa lepida
Suaeda esteroa                                 x       x
Suaeda nigra
Suaeda taxifolia
Triglochin maritima                    x       x       x
Vulpia microstachys var.
  microstachys

                                        Low          High
Scientific name                      transition   transition   Grass

Abronia maritima                                      x
Abronia umbel lata                                    x
Achillea millefolium                                  x          x
Acmispon glaber                                       x          x
Ambrosia psilostachya
Artemisia californica
Artemisia douglasiana                    x            x
Artemisia tridentata                                             x
Arthrocnemum subterminale
Astragalus pycnostachyus var.            x
  lanosissimus
Astragalus tener var. titi               x            x
Atriplex californica                     x            x
A trip lex canescens
Atriplex lentiformis
Atriplex prostrata                       x
Atriplex watsonii                        x            x
Baccharis pilularis                                   x
Baccharis salicifolia subs,
  salicifolia
Baccharis sarothroides
Batis maritima                           x
Cressa truxillensis
Croton califomicus                                    x          x
Distichlis littoralis
Distichlis spicata                       x
Encelia californica                                              x
Eriogonum fasciculatum
Frankenia salina                         x            x
Grindelia camporum
Hazardia squarrosa                                               x
Heteromeles arbutifolia                                          x
Hordeum brachyantherum                   x            x
Isocoma menziesii                                     x          x
Iva axillaris
Jaumea camosa
Juncus acutus subs, leopoldii
Juncus bufonius                          x
Limonium californicum                    x
Lupinus chamissonis
Lycium californicum                      x            x
Melica imperfecta                                                x
Mimulus aurantiacus                                              x
Oenothera elata                                                  x
Peritoma arborea
Phacelia ramosissima                                  x          x
Plantago erect a                                                 x
Platanus racemosa
Populus fremontii subs, fremontii
Potentilla anserina subs, pacifwa        x            x
Pseudognaphalium californicum
Rosa californica                                      x          x
Salicomia bigelovii
Salicomia pacifwa
Salix exigua subs, exigua
Salix lasiolepis
Salvia apiana
Salvia mellifera
Schoenoplectus acutus var.
  occidentalis
Schoenoplectus califomicus
Spartina foliosa
Stipa cemua                                                      x
Stipa lepida                                                     x
Suaeda esteroa
Suaeda nigra
Suaeda taxifolia
Triglochin maritima
Vulpia microstachys var.                                         x
  microstachys

                                             Fresh     Salt
Scientific name                      Scrub   water   tolerant

Abronia maritima                                        x
Abronia umbel lata                                      x
Achillea millefolium                                    x
Acmispon glaber                        x                x
Ambrosia psilostachya
Artemisia californica                  x
Artemisia douglasiana                                   x
Artemisia tridentata                   x                x
Arthrocnemum subterminale                               x
Astragalus pycnostachyus var.                           x
  lanosissimus
Astragalus tener var. titi             x                x
Atriplex californica                   x                x
A trip lex canescens                   x                x
Atriplex lentiformis                   x                x
Atriplex prostrata                                      x
Atriplex watsonii                                       x
Baccharis pilularis                    x
Baccharis salicifolia subs,            x
  salicifolia
Baccharis sarothroides                 x                x
Batis maritima                                          x
Cressa truxillensis
Croton califomicus                     x                x
Distichlis littoralis                                   x
Distichlis spicata                                      x
Encelia californica                    x
Eriogonum fasciculatum                 x
Frankenia salina                                        x
Grindelia camporum                     x                x
Hazardia squarrosa                     x                x
Heteromeles arbutifolia                x       x
Hordeum brachyantherum                         x        x
Isocoma menziesii                      x                x
Iva axillaris
Jaumea camosa                                           x
Juncus acutus subs, leopoldii
Juncus bufonius                                         x
Limonium californicum                                   x
Lupinus chamissonis                    x
Lycium californicum                                     x
Melica imperfecta                      x
Mimulus aurantiacus                    x
Oenothera elata                                         x
Peritoma arborea                       x                x
Phacelia ramosissima                   x                x
Plantago erect a                       x                x
Platanus racemosa                      x       x
Populus fremontii subs, fremontii      x       x
Potentilla anserina subs, pacifwa                       x
Pseudognaphalium californicum
Rosa californica                       x                x
Salicomia bigelovii                                     x
Salicomia pacifwa                                       x
Salix exigua subs, exigua                      x
Salix lasiolepis                               x
Salvia apiana                          x                x
Salvia mellifera                       x                x
Schoenoplectus acutus var.                     x
  occidentalis
Schoenoplectus califomicus                     x        x
Spartina foliosa                                        x
Stipa cemua                            x
Stipa lepida                           x
Suaeda esteroa                                          x
Suaeda nigra                                            x
Suaeda taxifolia
Triglochin maritima                                     x
Vulpia microstachys var.               x                x
  microstachys

Appendix II. Detailed, species-specific seed collection, seed
germination, and seed storage information for wetland, coastal sage
scrub, and upland transition species native to south-ern California.
Information is sorted alphabetically by scientific name.

                                                    Fruit/seed
Scientific name             Start    End          characteristics

Abronia maritima             May     Aug     Winged fruit 10-14 mm
  (Drennan 2008, Baldwin                       long. Fruit contains
  et al. 2012)                                 single-seeded achenes.
Abronia umbellate            May     Aug     Winged fruit 6-13 mm
  (Drennan 2008, Baldwin                       long. Fruit contains
  et al. 2012, Center                          single-seeded achenes.
  for Plant Conservation
  2015,)
Achillea millefolium         Aug     Oct     Oblong fruit, usually 2
  (Baskin and Baskin                           mm in length, contains
  2002a, Baldwin                               brown disk achenes.
  et al. 2012)                                 Seeds mature in late
                                               summer-early fall.
Acmispon glaber              May     Jul     Narrow, bean-shaped,
  (Montalvo and Beyers                         curved seed pods 1-2
  2010c)                                       mm long. Indehiscent
                                               pods ripen in 4-6
                                               weeks. Mature pods are
                                               dry and brown or olive
                                               green.
Ambrosia psilostachya        Oct     Dec     Brown bur 3-4.5 mm long
  (Pavek 1992, Baldwin                         contains tiny achenes.
  et al. 2012)
Artemisia californica        Oct     Feb     Fruit 0.8-1.5 mm long.
  (Hauser 2006, Young-                         Very small achenes,
  Mathews 2010, Baldwin                        generally mature in
  et al. 2012)                                 early fall or winter;
                                               wind dispersed.
Artemisia douglasiana                        <1 mm, glabrous fruit.
  Elkhom Slough National                       Small, ellipsoid,
  Estuarine Research                           hairless achenes
  Reserve 2001,                                without ribs or
  Shultz 2014)                                 angles.
Artemisia tridentate         Sep    early    Fruit glandular or
  (Elkhom Slough                    winter     hairy, 1-2 mm in
  National Estuarine                           length. Very small
  Research Reserve 2001,                       achenes, generally
  Baldwin et al. 2012,                         mature in early fall
  Tilley et al., n.d.)                         or winter.
Arthrocnemum                 Oct     Dec     Stems have tiny flowers
  subterminale (Zedler                         that occur below the
  2001, Clarke et al.                          tip of the stem and
  2007, Baldwin et al.                         contain brown,
  2012)                                        vertical seeds with
                                               hard seed coat, 1-1.4
                                               mm in length.
Astragalus pycnostachyus     Jul             Fruit ovate and
  var. ianosissimus                            inflated; 6-11 mm long
  (McCue 2010, Baldwin                         and 3.5-6 mm wide.
  et al. 2012, U.S. Fish                       Seeds are smooth,
  and Wildlife Service,                        compressed with a
  n.d.)                                        small notch at
                                               attachment site. 2 or
                                               more seeds/fruit.
Astragalus tetter var.       May             Fruit 6-50 mm long and
  titi (Showers 2010,                          1.7-3.5 mm wide. Seeds
  Baldwin et al. 2012)                         are smooth, compressed
                                               with a small notch at
                                               attachment site. 2 or
                                               more seeds/fruit.
Atriplex calijarnica         Sep     Oct     Mature fruit is an
  (Young 2001a)                                utricle with 1 seed.
                                               Seeds are black,
                                               shiny, hard, round,
                                               and flat; 2 mm at
                                               maturity.
Atriplex canescens           Oct     Apr     Cream-colored 4 winged
  (Springfield 1970,                           utricle, 5-23 mm wide.
  Baldwin et al. 2012)                         Seeds 1.5-2.5 mm long
                                               with brown, papery
                                               inner seed coat.
                                               Species has high
                                               percentage of empty
                                               seed. Smaller fruits
                                               tend to have higher
                                               percentages of filled
                                               seed.
Atriplex lentiformis         Sep     Jan     Produces large amounts
  (Young et al. 1980,                          of dark brown, 1.5 mm
  Baldwin et al. 2012)                         long seeds.
Atriplex prostrated          Sep     Oct     Two types of seeds:
  (Khan and Ungar 1984,                        brown, 1-2.5 mm long,
  Zedler 2001, Baldwin                         and black, 1-1.5 mm
  et al. 2012)                                 long.
Atriplex watsonii            Jun     Sep     Seeds light brown, about
  (Zedler 2001,                                1 mm long.
  Bryant 2004)
Baccharis pilularis          Aug     Dec     Single-sex white
  (Bonner and Karrfalt                         flowers. Glabrous,
  2008, Montalvo et al.                        ribbed fruit 1-2 mm
  2010b, Baldwin et al.                        long, pappus 5.5-9 mm
  2012)                                        long. Mature seeds are
                                               tiny, dark brown
                                               achenes with ring of
                                               long, unbranched
                                               pappus bristles.
Baccharis salicifolia        May     Jul     Glabrous, ribbed fruit
  subs. Salicifolia                            0.8-1.3 mm long,
  (Bonner and Karrfalt                         pappus 3-6 mm long.
  2008, Baldwin et al.                         Tiny achenes with a
  2012)                                        bristly pappus.
Baccharis sarothroides                       Glabrous, ribbed fruit
  (Bonner and Karrfalt                         2-2.6 mm long, pappus
  2008, Baldwin et al.                         2-3 mm long. Tiny
  2012)                                        achenes.
Bads maritime                Oct     Nov     Hard-walled lenticular
  (Zedler 2001, Marcone                        or oblong seeds. 1 mm
  2003, Francis 2009,                          in length.
  Lonard et al. 2011)
Cressa truxtilensis          Jul     Aug     Fruits are small hairy
  (Elkhom Slough                               capsules, 1/8" long.
  National Estuarine                           Seeds pinkish in
  Research Reserve 2001,                       color, broadly
  Zedler 2001)                                 egg-shaped.
Croton californicus          Jul     Nov     Flowers develop into
  (Young 2001b, Baldwin                        compact, greenish seed
  et al. 2012)                                 pods. Mature seeds are
                                               smooth, round, and
                                               brown with tan spots.
                                               3.5-5.5 mm in length.
Distichlis lift oralis       Jun     Sep     Spikelets, 8-13 mm in
  (Zedler 2001, Clarke                         length, generally
  et al. 2007, Baldwin                         concealed by leaves.
  et al. 2012)                                 Seeds are quite small
                                               and remain in flowers
                                               until senescence.
Distichlis spicata           Sep     Nov     Spikelets 6-20 mm long.
  (Baskin and Baskin                           Seed likely dormant at
  2002b, Elsey-Quirk                           time of dispersal.
  et al. 2009, Baldwin
  et al. 2012)
Encelia californica                          Fruit 5-7 mm long,
  (Bonner and Karrfalt                         slightly longer with
  2008, Baldwin et al.                         pappus. Seeds dark
  2012)                                        brown at maturity.
Eriogonum fasciculatum       May     Aug     Glabrous fruit 1.8-2.5
  (Zedler2001, Montalvo                        mm in length.
  and Beyers 2010a,
  Baldwin et al. 2012)
Fran ken ia salina           Sep     Oct     Ellipsoid seed capsules
  (Young 2001k)                                (8 mm) contain 1 mm
                                               long, brownish black
                                               seeds. Ovular in shape
                                               with pointed tips.
Grindelia camporum           Jun     Oct     Small, long, and flat
  (Zafar and Shah 1994,                        achenes. Wind-borne,
  Bliss 2012)                                  dandelion-like achenes
                                               with feathery tufts.
Hazardia squarrosa                           Fruit: 5-8 mm, 5-angled,
  (Keil et al. 2013)                           glabrous; pappus 7-12
                                               mm, white to red-brown
                                               in color. Oblong to
                                               lanceolate seeds.
Heteromeles arbutifolia      Oct     Jan     Large, smooth brown
  (Bonner and Karrfalt                         seeds. 2-3 seeds per
  2008, Baldwin et al.                         pome.
  2012, Gordon 2014,
  Recon Native Plants
  Inc. 2015)
Hordeum brachyantherum       Jun     July    Mature inflorescences
  (Elkhom Slough                               are light brown.
  National Estuarine
  Research Reserve 2001,
  Young 2001d)
Isocoma menziesii            Sep     Nov     Tan-colored achenes,
  (Zedler 2001, Wall and                       longer than wide,
  Macdonald 2009,                              wider on the plumose
  Montalvo and Beyers                          end, with lengthwise
  2010b)                                       striations. The top of
                                               the achene has a ring
                                               of white bristles.
                                               Seeds mature when the
                                               pappus becomes fluffy
                                               and achenes detach
                                               easily from the
                                               receptacle.
Iva axillaris                                1-2 seeds/head. 2 mm
  (Montalvo et al.                             long, turnip-shaped,
  2010b)                                       light, and buoyant.
Jaumea carnosa               Jul     Oct     Seeds are linear achenes
  (Young 2001e)                                with longitudinal
                                               stripes.
Juncus acutus subs.          Aug     Nov     Shiny brown capsules
  leopoldii                                    contain multiple
  (Zedler2001, Baldwin                         irregularly shaped
  et al. 2012)                                 seeds. Seeds can be
                                               narrowly winged.
Juncus bufonius              Mar     May     Ovoid or elliptic seeds.
  (Zedler 2001, Baldwin                        Seeds generally
  et al. 2012)                                 0.3-0.6 mm long.
Limonium californicum        Sep     Nov     3 mm long narrow
  (Young 200If)                                ellipse, dark brown/
                                               red at maturity.
Lupinus chamissonis          Apr     Jun     Hairy legume pods
  (Young 200 lg)                               2.5-3.5 cm long.
                                               Mature seeds are dark
                                               brown and speckled and
                                               3-4 mm in length.
Lycium californicum          Jan     Feb     3-6 mm red berries. 2
  (Zedler 2001, Baldwin                        oblong seeds per
  et al. 2012)                                 berry.
Melica imperfecta            Apr     Jun     Mature inflorescences
  (Ashby and Hellmers                          are brown; mature
  1959, Emery 1988,                            seeds are tan.
  Young 2001h, Baskin
  and Baskin 2002c)
Mimulus aurantiacus          Jun     Aug     Mature capsules are
  (Young 2001c, Baldwin                        brown and contain
  et al. 2012)                                 tiny, black seeds less
                                               than 1 mm in length.
Oenothera elata                              Seeds are irregularly
  (Greiner and Kohl                            shaped, stacked in
  2014, B & T World                            small, brown, woody
  Seeds 2015, Dave's                           capsules with four
  Garden 2015a,                                chambers each with two
  Kleinman, n.d.)                              rows of small seeds.
Peritoma arborea                             Capsules 3-6 cm long and
  (Lippitt et al. 1994,                        1-2.5 cm wide. Mature
  Borders et al. 2008,                         fruits will often
  Baldwin et al. 2012)                         split at the seam,
                                               revealing seeds.
                                               Dark-colored seeds
                                               tend to be more viable
                                               that light-colored
                                               seeds.
Phacelia ramosissima                         Capsules contain 8-12,
  (Baldwin et al. 2012)                        1-2 mm long pitted
                                               seeds.
Plantago erecta              Apr             2-2.5 mm long.
  (Gulmon 1992, Montalvo
  et al. 2010a, Baldwin
  et al. 2012)
Platanus racemosa             Jun   Spring   Chestnut brown seed pods
  (Bonner and Karrfalt                         at maturity, many are
  2008)                                        empty. Achenes are
                                               2-2.5 mm in length and
                                               have small tuft at
                                               base.
Populus fremontii subs.      Mar     Aug     Capsules contain seeds
  fremontii (Gulmon 1992,                      with long, silky
  Stettler, 1996, Clarke                       hairs.
  et al. 2007, Kleinman,
  n.d.)
Potentilla anserina                          Fruits are oval, flat,
  subs. pacifica (Walker                       and reddish-brown and
  2005, Stevens, n. d.,                        about 2 mm in length.
  Baldwin et al. 2012)
Pseudognaphalium                             Oblong fruits with
  californicum                                 bristly, tuft-like
  (Keeley and Keeley                           projections (shed at
  1987, Nesom 2013)                            maturity).
Rosa californica             Jul     Sep     Mature fruits (rose
  (Young 2001i, Lady                           hips) are bright red.
  Bird Johnson                                 Each hip contains
  Wildflower Center                            multiple seeds.
  2007)
Salicornia bigelovii         Sep     Nov     1-1.5 mm curved, hairy
  (Glenn et al. 1997,                          seeds.
  Zedler 2001, Baldwin
  et al. 2012)
Salicornia pacifica          Oct     Nov     Mature seeds pinkish
  (Khan and Weber 1986,                        white, puberulent, and
  Young 200 lj)                                0.5-1 mm long.
                                               Sequentially
                                               hermaphroditic.
Salix exigua subs,           May     July    Glabrous ovular
  exigua (Young and                            capsules. Small seeds
  Clements 2003,                               with pappus. Normally
  Anderson 2006, Clarke                        dispersed via wind or
  et al. 2007)                                 water.
Salix lasiolepis             May             Glabrous ovular
  (Bonner and Karrfalt                         capsules.
  2008, Don 2014)

Salvia apiana               July     Aug     Shiny, light brown
  (Stevens 1994,                               fruit. Fruits are
  Montalvo and Beyers                          2.5-3 mm in length.
  2010d, Baldwin et al.
  2012, Native Plant
  Database 2015)
Salvia mellifera             Jun     Aug     Dry calyces are gravity
  (Montalvo and Beyers                         dispersed. Up to 4
  2010e)                                       seeds/calyx. Seeds are
                                               1 mm by 2 mm.
Sclioenoplectus acutus       Aug     Sep     Wide, smooth fruits with
  var. occidentalis                            2 or 3 distinct sides.
  (Lacroix and Mosher                          Fruit 2-3 mm long and
  1995, Johnson 2004,                          1.2-1.7 mm wide.
  Baldwin et al. 2012,
  Baskin and Baskin
  2014)
Schoenoplectus                               2-sided, smooth fruits.
  californicus                                 Fruits 1.8-2.2 mm long
  (Stevens 2003,                               and 1.3 mm wide.
  Baldwin et al. 2012)
Spartina foliosa             Sep     Nov     10-25 mm spikelets.
  (Zedler 2001, Baldwin
  et al. 2012)
Stipa cernua                 Jul     Aug     Linear, smooth, glabrous
  (Laude et al. 1952,                          seed.
  Amme 2003, Herrera
  et al. 2006)
Stipa lepida                                 Brown fruit. Dark seeds
  (Ashby and Hellmers                          4-7 mm in length.
  1959, Elkhom Slough
  National Estuarine
  Research Reserve 2001,
  Amme 2003, Dave's
  Garden 2015b)
Suaeda esteroa               Oct     Dec     Two types of seeds:
  (Zedler 2001, Baldwin                        seeds can be
  et al. 2012)                                 lenticular, black, and
                                               shiny (0.8-1.7 mm in
                                               length) or horizontal
                                               and matte (1-1.5 mm in
                                               length).
Suaeda nigra                 Sep     Oct     Small, lenticular, shiny
  (Borders, n.d.)                              black seeds 0.5-2 mm
                                               long. Seed coat can be
                                               smooth, finely dotted,
                                               warty, net-like, or
                                               prickly.
Suaeda taxifolia             Jun     Jul     1-2 mm horizontal or
  (Zedler 2001, Baldwin                        vertical seeds. Seeds
  et al. 2012)                                 are shiny, lenticular
                                               and range from black
                                               to brown.
Triglochiit maritima         Jul     Sep     Mature inflorescences
  (Young 2002, Baldwin                         are brown. 1 seed/
  et al. 2012, Recon                           fruit. Seeds of genus
  Native Plants Inc.                           usually linear. Seeds
  2015)                                        can be flat or angled.
Vulpia microstachys                          5.5-10 mm spikelets
  (Young and Young 1986,                       contain 4-6 mm long
  Howard 2006, Baldwin                         fruits.
  et al. 2012)

Scientific name                    Seed collection details

Abronia maritima            Plants seed throughout the year,
  (Drennan 2008, Baldwin      majority of seed production
  et al. 2012)                occurs in late spring/summer.
                              Removal of woody capsules
                              aided with the use of generic
                              nail clippers.
Abronia umbellate           Plants seed throughout the year,
  (Drennan 2008, Baldwin      majority of seed production
  et al. 2012, Center         occurs in late spring/summer.
  for Plant Conservation      Removal of woody capsules
  2015,)                      aided with the use of generic
                              nail clippers.
Achillea millefolium        Cut entire inflorescences, collect
  (Baskin and Baskin          in paper bags. Clean seeds with
  2002a, Baldwin              a hammermill, screen, and
  et al. 2012)                fanning mill.
Acmispon glaber             Strip ripe seed pods from stems
  (Montalvo and Beyers        by hand. Avoid breaking seeds
  2010c)                      during thrashing. Rub pods
                              with wooden block over #16
                              (medium) screen. Remove
                              seeds from pods. Remove
                              excess chaff with seed blower.
Ambrosia psilostachya
  (Pavek 1992, Baldwin
  et al. 2012)
Artemisia californica       Strip brown inflorescence
  (Hauser 2006, Young-        by hand.
  Mathews 2010, Baldwin
  et al. 2012)
Artemisia douglasiana       Seed is ready to harvest when it
  Elkhom Slough National      can be easily removed from the
  Estuarine Research          heads by shaking. Clip seed
  Reserve 2001,               stalks and air dry in a paper
  Shultz 2014)                bag. To thresh seeds rub the
                              inflorescence through a screen.
                              Remove chaff with a blower.
Artemisia tridentate        Seed from genus is ready to
  (Elkhom Slough              harvest when it can be easily
  National Estuarine          removed from the heads by
  Research Reserve 2001,      shaking. Clip seed stalks and
  Baldwin et al. 2012,        air dry in a paper bag. To thresh
  Tilley et al., n.d.)        seeds mb the inflorescence
                              through a screen. Remove chaff
                              with a blower.
Arthrocnemum                Best to collect in November.
  subterminale (Zedler        Collect inflorescences and air
  2001, Clarke et al.         dry. When dry, shake seeds
  2007, Baldwin et al.        from stalks.
  2012)
Astragalus pycnostachyus    Other plants in genus, specifically
  var. ianosissimus           A. sinuatus, have seeds that
  (McCue 2010, Baldwin        mature in late July.
  et al. 2012, U.S. Fish
  and Wildlife Service,
  n.d.)
Astragalus tetter var.      Endangered species. Extract seeds
  titi (Showers 2010,         from fruits by hand. Thresh seeds
  Baldwin et al. 2012)        over sieve large enough to let set
                              seed pass through. Run seeds
                              through seed blower to remove
                              parasitized or aborted seed.
Atriplex calijarnica        Gently seeds rub over # 18 sieve.
  (Young 2001a)               Remove as much chaff as
                              possible with a seed blower.
Atriplex canescens          Strip seeds from branches by
  (Springfield 1970,          hand. If available, use a
  Baldwin et al. 2012)        hammermill and a fanning mill
                              to de-wing and clean seed.
                              Collections made later (Dec-Apr)
                              tend to have higher germination
                              rates.
Atriplex lentiformis        Dioecious.
  (Young et al. 1980,
  Baldwin et al. 2012)
Atriplex prostrated         Fully mature fruit can be shaken
  (Khan and Ungar 1984,       or hand stripped from
  Zedler 2001, Baldwin        branches. Seeds will often
  et al. 2012)                remain on bushes until April,
                              so late collections are possible.
Atriplex watsonii
  (Zedler 2001,
  Bryant 2004)
Baccharis pilularis         Dioecious. Collect seed heads by
  (Bonner and Karrfalt        hand into open breathable bags.
  2008, Montalvo et al.       Alternatively shake branches over
  2010b, Baldwin et al.       a tarp. Fruit should be spread out
  2012)                       to dry in a well-ventilated room or
                              in the sun. Rub dried heads
                              between palms or over a screen to
                              remove the pappus and phyllaries.
Baccharis salicifolia       Dioecious. Collect ripe fruits by
  subs. Salicifolia           hand or by shaking seeds onto
  (Bonner and Karrfalt        canvases/tarps. Dry seeds at
  2008, Baldwin et al.        room temperature. Once dry,
  2012)                       rub seeds over a screen to
                              remove the pappus.
Baccharis sarothroides      Dioecious. Seeds can be collected
  (Bonner and Karrfalt        by hand or branches can be
  2008, Baldwin et al.        shaken above tubs/tarps.
  2012)
Bads maritime               Collect when fruits mature and
  (Zedler 2001, Marcone       turn from green to white.
  2003, Francis 2009,         Extract seed from fruit. Dried
  Lonard et al. 2011)         fruits should fragment easily,
                              exposing seed
Cressa truxtilensis         Produces mature seeds from late
  (Elkhom Slough              summer into early autumn.
  National Estuarine
  Research Reserve 2001,
  Zedler 2001)
Croton californicus         Dioecious. Collect July 15th-
  (Young 2001b, Baldwin       November 17th. Remove chaff
  et al. 2012)                by hand. Remove seeds
                              from pods.
Distichlis lift oralis      Dioecious. Strip flowers with
  (Zedler 2001, Clarke        seeds from inflorescences.
  et al. 2007, Baldwin
  et al. 2012)
Distichlis spicata          Dioecious. Seed is 2 mm long
  (Baskin and Baskin          and brownish-gray at maturity.
  2002b, Elsey-Quirk          Rub seeds over #18 sieve to
  et al. 2009, Baldwin        clean.
  et al. 2012)
Encelia californica         Achenes are wedge-shaped and
  (Bonner and Karrfalt        densely compressed. Edges are
  2008, Baldwin et al.        long-ciliate and faces are
  2012)                       glabrous or short-hairy.
                              Collection timing is critical as
                              achenes are easily blown from
                              plant after reaching maturity.
Eriogonum fasciculatum      Best to collect from Jun-Jul.
  (Zedler2001, Montalvo       Collect inflorescences as they
  and Beyers 2010a,           begin to turn rusty brown. Push
  Baldwin et al. 2012)        seeds through a screen to
                              remove chaff.
Fran ken ia salina          Collect: September 16th- October
  (Young 2001k)               21st. Collect mature flowers
                              and rub over #25 sieve. Use
                              gloves when handling, the
                              plant can be spiky.
Grindelia camporum          Harvest seed in June and again in
  (Zafar and Shah 1994,       October. Clip seed heads or
  Bliss 2012)                 shake/rub mature seeds from
                              seed heads into a collection
                              bag. To clean, rub seed heads
                              over sieve. Remove chaff using
                              additional sieves or an air
                              separator. Air dry in oven at
                              203 [degrees]F.
Hazardia squarrosa
  (Keil et al. 2013)
Heteromeles arbutifolia     Clip or strip fruits from branches
  (Bonner and Karrfalt        when bright red. Soak berries in
  2008, Baldwin et al.        water to ferment (over-soaking
  2012, Gordon 2014,          can be damaging). Pulp should
  Recon Native Plants         float, making it easier to separate
  Inc. 2015)                  seeds from pulp. Alternatively,
                              pulse berries in blender and then
                              mb mixture over a screen to
                              isolate fruit. Dry seeds before
                              storing. RECON suggests
                              keeping fruit intact.
Hordeum brachyantherum      Seed easily removed when stalks
  (Elkhom Slough              are hand stripped. No
  National Estuarine          additional cleaning required.
  Research Reserve 2001,
  Young 2001d)
Isocoma menziesii           Collect achenes golden in color, as
  (Zedler 2001, Wall and      seeds are usually eaten by time
  Macdonald 2009,             achenes turn brown. Shake ripe
  Montalvo and Beyers         heads over open containers to
  2010b)                      collect achenes. Alternatively,
                              remove ripe heads and keep in
                              porous bags. For 7. acradenia,
                              Wall and Macdonald recommend
                              rubbing flowers over a large
                              screen, using a seed blower, and
                              sieving over a #18 screen to
                              separate seeds from bracts.
Iva axillaris               Strip seeds by hand or beat into a
  (Montalvo et al.            hopper/open container. Rub
  2010b)                      flower material over a screen
                              and run through a blower to
                              remove chaff.
Jaumea carnosa              Collect seed while fruits are
  (Young 2001e)               swollen and green. Rub seeds
                              over # 12 sieve to clean.
Juncus acutus subs.
  leopoldii
  (Zedler2001, Baldwin
  et al. 2012)
Juncus bufonius             Seed capsules dehisce; seeds
  (Zedler 2001, Baldwin       should be collected quickly
  et al. 2012)                after plant death. Shake mature
                              flowers to collect tiny seed.
Limonium californicum       Collections made in October are
  (Young 200If)               best. Collect entire flower
                              heads, which should detach
                              easily when ripe. Rub flower
                              heads over #20 sieve.
Lupinus chamissonis         Remove seeds from receptacles,
  (Young 200 lg)              no further cleaning required.
Lycium californicum         Pick by hand. Berries best
  (Zedler 2001, Baldwin       collected within 2 weeks of
  et al. 2012)                appearance, otherwise birds
                              will eat majority. Extract seeds
                              from berries within a week,
                              before berries begin to mold.
Melica imperfecta           Strip inflorescences.
  (Ashby and Hellmers
  1959, Emery 1988,
  Young 2001h, Baskin
  and Baskin 2002c)
Mimulus aurantiacus         Rub seed capsules over a sieve.
  (Young 2001c, Baldwin
  et al. 2012)
Oenothera elata             Collect seed from spring cultivars
  (Greiner and Kohl           in October and from winter
  2014, B & T World           cultivars in September. Bag
  Seeds 2015, Dave's          seed heads and allow them to
  Garden 2015a,               dry on plant or collect early and
  Kleinman, n.d.)             allow to ripen in paper bags.
Peritoma arborea            Flowers several times/year (except
  (Lippitt et al. 1994,       Dec-Jan). Ready for collection
  Borders et al. 2008,        when capsules turn brown and
  Baldwin et al. 2012)        are crisp. Strip mature fruits from
                              plants by hand. Break apart pods
                              by hand or with a hammermill or
                              coater blender.
Phacelia ramosissima        Collect seed when flowers are dry
  (Baldwin et al. 2012)       and brown. Strip seed from
                              mature inflorescences directly
                              into collection bag.
Plantago erecta             Dehiscent, ballistic seed dispersal.
  (Gulmon 1992, Montalvo      Collect inflorescences into a
  et al. 2010a, Baldwin       paper bag and let dry. Use sieve
  et al. 2012)                to clean.
Platanus racemosa           Collect seedpods after they have
  (Bonner and Karrfalt        turned brown. The task is
  2008)                       easiest after leaves have fallen.
                              Seedpods remain on trees into
                              spring. Cut seedpods directly
                              from tree. Crush dried
                              seedpods to open. Remove dust
                              and fine hairs.
Populus fremontii subs.     Dioecious. Collect seed as it is
  fremontii (Gulmon 1992,     released from capsules during
  Stettler, 1996, Clarke      dehiscence or collect entire
  et al. 2007, Kleinman,      catkins prior to dehiscence.
  n.d.)                       Separate cotton fibers
                              from seed.
Potentilla anserina         Let seeds dry on plant prior to
  subs. pacifica (Walker      collection.
  2005, Stevens, n. d.,
  Baldwin et al. 2012)
Pseudognaphalium
  californicum
  (Keeley and Keeley
  1987, Nesom 2013)
Rosa californica            Collect hips as soon as they are
  (Young 2001i, Lady          ripe. Extract seeds by hand from
  Bird Johnson                dried hips. Alternatively,
  Wildflower Center           macerate hips in water; remove
  2007)                       floating seeds.
Salicornia bigelovii        Entire inflorescences should be
  (Glenn et al. 1997,         collected and air-dried. When
  Zedler 2001, Baldwin        dry, strip seeds from
  et al. 2012)                inflorescences.
Salicornia pacifica         Collect inflorescences when plant
  (Khan and Weber 1986,       tips are purple. Dry seeds on a
  Young 200 lj)               screen for up to 3 months.

Salix exigua subs,          Dioecious. Harvest when catkins
  exigua (Young and           are yellow-brown and capsules
  Clements 2003,              begin to open. Shake catkins to
  Anderson 2006, Clarke       remove dried seeds.
  et al. 2007)
Salix lasiolepis            Dioecious. Hand-harvest catkins
  (Bonner and Karrfalt        when they begin to turn yellow/
  2008, Don 2014)             brown. It is recommended to
                              wait until capsules open.
                              Separate seeds from cotton.
Salvia apiana               Collect seeds as capsules begin to
  (Stevens 1994,              dry, before seeds are dispersed.
  Montalvo and Beyers         Shake seeds from seed heads.
  2010d, Baldwin et al.       Use a sieve to isolate seeds.
  2012, Native Plant
  Database 2015)
Salvia mellifera            Collect after inflorescences with
  (Montalvo and Beyers        calyces are dry and brown.
  2010e)                      Collect mature seeds by clipping,
                              stripping, or shaking seed heads.
                              Seed should be dried and passed
                              through a sieve. Use of a blower
                              is recommended.
Sclioenoplectus acutus      Because they are easily dispersed
  var. occidentalis           by wind, it is important to
  (Lacroix and Mosher         collect seeds close to the time
  1995, Johnson 2004,         of maturity. Seeds must be
  Baldwin et al. 2012,        separated from the panicle and
  Baskin and Baskin           cleaned.
  2014)
Schoenoplectus              Harvest seed by hand from seed
  californicus                heads. Alternatively, use shears
  (Stevens 2003,              to clip entire seed heads from
  Baldwin et al. 2012)        plant. Clean seeds.
Spartina foliosa            Multiple harvests may increase
  (Zedler 2001, Baldwin       probability of collected good
  et al. 2012)                seeds prior to dispersal or
                              herbivory loss.
Stipa cernua                Harvest by hand or with a flow-vac
  (Laude et al. 1952,         or combine at maturity. Collection
  Amme 2003, Herrera          possible for 2-3 weeks.
  et al. 2006)
Stipa lepida                Seeds mature in spring. Allow
  (Ashby and Hellmers         seed to mature on plant. At
  1959, Elkhom Slough         maturity, harvest seed. Clean
  National Estuarine          prior to storage.
  Research Reserve 2001,
  Amme 2003, Dave's
  Garden 2015b)
Suaeda esteroa              Best to collect in Nov or early
  (Zedler 2001, Baldwin       Dec. Cut whole inflorescence
  et al. 2012)                or strip inflorescence. After
                              cleaning, seeds should be dried.
Suaeda nigra                Collect when seeds are hard,
  (Borders, n.d.)             black, and shiny when calyces
                              will be brown and crumbly.
                              Strip seeds from stalk by hand.
                              Pass seed through a hammer
                              mill or a sieve prior. Seeds
                              should be spread out to dry
                              before being processed/stored.
Suaeda taxifolia            Strip inflorescence by hand.
  (Zedler 2001, Baldwin
  et al. 2012)
Triglochiit maritima        Collect seeds between July 17-Sept.
  (Young 2002, Baldwin        23rd. Rub dry fruits between
  et al. 2012, Recon          fingers to extract the seeds.
  Native Plants Inc.
  2015)
Vulpia microstachys         Unknown exactly when seeds
  (Young and Young 1986,      from S. California plants
  Howard 2006, Baldwin        mature (intermountain varieties
  et al. 2012)                mature late July-late September).

                                                            Longevity
Scientific name                   Seed germination            (yrs.)

Abronia maritima            In the most successful              3
  (Drennan 2008, Baldwin      trials, achenes removed
  et al. 2012)                from anthrocarp. Place
                              achenes on filter paper
                              in sterile petri dishes
                              with ethephon or other
                              10-100 umol ethylene
                              source. Incubate achenes
                              in a chamber with
                              alternating 12 h periods
                              of light (27[degrees]C)
                              and dark (20[degrees]C).
                              Requires a sandy
                              substrate.
Abronia umbellate           In the most successful              3
  (Drennan 2008, Baldwin      trials, achenes removed
  et al. 2012, Center         from anthrocarp. Some seed
  for Plant Conservation      lots require cold
  2015,)                      pre-treatment. Germination
                              requirements may differ
                              year to year. For best
                              results, sow clean seeds
                              in the top 1" of a sandy
                              growing medium.
Achillea millefolium        Lightly cover seeds with           3-5
  (Baskin and Baskin          growing medium (milled
  2002a, Baldwin              sphagnum, peat, perlite,
  et al. 2012)                vermiculite w/ oscmocote).
                              90-100% germination rate.
Acmispon glaber             Heat or mechanical              Long-lived
  (Montalvo and Beyers        scarification needed to
  2010c)                      break dormancy. Soak seeds
                              in boiling water or heat
                              in 120[degrees]C oven for
                              5 minutes for highest
                              yield.
Ambrosia psilostachya                                          3-5
  (Pavek 1992, Baldwin
  et al. 2012)
Artemisia californica       Seeds will germinate when          2-5
  (Hauser 2006, Young-        fresh. Stored seeds need
  Mathews 2010, Baldwin       to be exposed to light and
  et al. 2012)                may require cold
                              stratification. Other
                              sources imply that
                              pre-germination treatment
                              is not necessary.
Artemisia douglasiana       Germinates naturally at            2-5
  Elkhom Slough National      relatively cool temps.
  Estuarine Research
  Reserve 2001,
  Shultz 2014)
Artemisia tridentate        No pre-germination treatment       2-5
  (Elkhom Slough              necessary.
  National Estuarine
  Research Reserve 2001,
  Baldwin et al. 2012,
  Tilley et al., n.d.)
Arthrocnemum                Seeds are highly germinable.       3-5
  subterminale (Zedler        Germination promoted by
  2001, Clarke et al.         low salinities.
  2007, Baldwin et al.
  2012)
Astragalus pycnostachyus    Flard seed coat may require     Long-lived
  var. ianosissimus           scarification.
  (McCue 2010, Baldwin
  et al. 2012, U.S. Fish
  and Wildlife Service,
  n.d.)
Astragalus tetter var.      If stored for an extended       Long-lived
  titi (Showers 2010,         period of time, hard seed
  Baldwin et al. 2012)        coat may require
                              scarification to initiate
                              germination. 95%
                              germination success rate
                              on 0.5% agar plates with
                              11 hours light at
                              20[degrees]C and 13 hours
                              dark at 12[degrees]C.
Atriplex calijarnica        Pre-planting: soak in water         10
  (Young 2001a)               for 24 hours, rinse. 86%
                              germination rate after
                              sowing in peat moss,
                              perlite, nutrients,
                              gypsum, and dolomitic
                              lime. Germination occurs
                              after 10 days.
Atriplex canescens          Germination is inhibited by        5-7
  (Springfield 1970,          lack of aeration, but
  Baldwin et al. 2012)        improved with de-winging.
                              Sow in medium with high
                              substrate moisture at low
                              temperatures, ideally at
                              18-24[degrees]C in
                              California. Early
                              collections may benefit
                              from a 60 min. soak in
                              sulfuric acid or a
                              pre-chill at 5[degrees]C
                              for 12 weeks.
Atriplex lentiformis        Maximum germination between        3-6
  (Young et al. 1980,         10-25[degrees]C.
  Baldwin et al. 2012)
Atriplex prostrated         Readily propagated from             10
  (Khan and Ungar 1984,       seed. In the field,
  Zedler 2001, Baldwin        germinates in late spring.
  et al. 2012)
Atriplex watsonii           Readily propagated from             10
  (Zedler 2001,               seed, germinates and
  Bryant 2004)                establishes easily in
                              field.
Baccharis pilularis         Seeds germinate without             1
  (Bonner and Karrfalt        pre-treatment. Cool
  2008, Montalvo et al.       temperatures yield highest
  2010b, Baldwin et al.       germination percentage.
  2012)
Baccharis salicifolia       No pre-treatment necessary.         1
  subs. Salicifolia           Light necessary for
  (Bonner and Karrfalt        germination.
  2008, Baldwin et al.
  2012)
Baccharis sarothroides      Germinates well in wet              1
  (Bonner and Karrfalt        soils.
  2008, Baldwin et al.
  2012)
Bads maritime               Difficult to grow from seed.        2+
  (Zedler 2001, Marcone       Marcone had success
  2003, Francis 2009,         exposing seeds to natural
  Lonard et al. 2011)         light conditions and using
                              a nutrient-enhanced
                              potting medium. No known
                              dormancy requirements.
Cressa truxtilensis                                             2
  (Elkhom Slough
  National Estuarine
  Research Reserve 2001,
  Zedler 2001)
Croton californicus         Pre-planting: soak seeds for        3
  (Young 2001b, Baldwin       24 hours in water, cold
  et al. 2012)                stratify for 30 days.
                              Should germinate 30 days
                              after sowing.
Distichlis lift oralis                                         4-5
  (Zedler 2001, Clarke
  et al. 2007, Baldwin
  et al. 2012)
Distichlis spicata          Seed germination highest           4-5
  (Baskin and Baskin          after wet stratification/
  2002b, Elsey-Quirk          a fluctuating inundation
  et al. 2009, Baldwin        regime and with low
  et al. 2012)                salinity (Elsey-Quirk,
                              2009). Soak in water for
                              24 hours before sowing.
                              Establishes well at
                              restoration sites.
Encelia californica         To break dormancy, pre-soak        2-5
  (Bonner and Karrfalt        seeds in water.
  2008, Baldwin et al.
  2012)
Eriogonum fasciculatum      Seeds germinate well in            2-5
  (Zedler2001, Montalvo       flats. Light improves
  and Beyers 2010a,           germination rate. Sow in
  Baldwin et al. 2012)        fall-early winter.
Fran ken ia salina          Seeds need no pretreatment.        0-2
  (Young 2001k)               Germination naturally
                              promoted by low salinity
                              and high temperatures in
                              spring.
Grindelia camporum          Soak in water under                3-5
  (Zafar and Shah 1994,       continuous light OR use
  Bliss 2012)                 two-stage cold
                              stratification at
                              32[degrees]F and
                              59[degrees]F in the dark
                              (Zafar 1994) to pre-treat.
Hazardia squarrosa                                             0-1
  (Keil et al. 2013)
Heteromeles arbutifolia     Fresh seeds germinate               2
  (Bonner and Karrfalt        readily. Chill stored
  2008, Baldwin et al.        seeds for 3 months at
  2012, Gordon 2014,          3-5[degrees]C prior to
  Recon Native Plants         sowing. Seeds germinate
  Inc. 2015)                  well 23[degrees]C.
Hordeum brachyantherum      No pre-treatment required.         4-5
  (Elkhom Slough              Sow seeds in May. Seeds
  National Estuarine          should germinate 21 days
  Research Reserve 2001,      after sowing. Germination
  Young 2001d)                rate: 60%.
Isocoma menziesii           No pre-treatment required.          1+
  (Zedler 2001, Wall and      Seeds germinate well in
  Macdonald 2009,             flats.
  Montalvo and Beyers
  2010b)
Iva axillaris               Generally exhibits low          Short-
  (Montalvo et al.            germination rates.              lived
  2010b)                      Scarification is not
                              effective. Cold
                              stratification may be
                              (studies needed).
Jaumea carnosa              Seeds germinate readily in          1
  (Young 2001e)               moist soil.
Juncus acutus subs.         Grows readily from seed in         2-5
  leopoldii                   moderate salinities.
  (Zedler2001, Baldwin        Clones can be dug entire
  et al. 2012)                and transplanted.
Juncus bufonius             Seeds germinate readily in         2-5
  (Zedler 2001, Baldwin       low salinity soils.
  et al. 2012)
Limonium californicum       Sow in April. Propagates            1
  (Young 200If)               readily from seeds or
                              plugs.
Lupinus chamissonis         Scarify using sandpaper for     Long-lived
  (Young 200 lg)              5 minutes. Then, soak in
                              hot water over night
                              (repeat for seeds that do
                              not imbibe). Sow in
                              growing medium
                              mid-October. Should
                              germinate after 3 days.
Lycium californicum         Soak seeds in water for at          1
  (Zedler 2001, Baldwin       least 12 hours, then
  et al. 2012)                transferred to moist
                              soil. Reported germination
                              rates are low, 5-10%.
Melica imperfecta           Plant has irregular                4-5
  (Ashby and Hellmers         germination patterns and
  1959, Emery 1988,           a low documented
  Young 2001h, Baskin         germination rate, 30%.
  and Baskin 2002c)           Literature inconsistent,
                              certain sources suggest
                              soaking seeds overnight in
                              fresh water and cold
                              stratifying for 2 weeks in
                              peat. Conversely, Emery
                              1988 feels no
                              pre-treatment necessary.
                              Constant light (i.e.
                              24-hour photoperiod)
                              exposure is documented to
                              speed flowering.
Mimulus aurantiacus         Sow seeds in August. No            2-5
  (Young 2001c, Baldwin       pre-treatment needed, 50%
  et al. 2012)                germination rate.
Oenothera elata             Surface sow (1 mm deep) to         3-5
  (Greiner and Kohl           ensure sufficient light.
  2014, B & T World           Long-day conditions (16
  Seeds 2015, Dave's          hours of light/ 8 hours of
  Garden 2015a,               darkness) should be
  Kleinman, n.d.)             simulated in the
                              greenhouse. Should
                              germinate after 15-30
                              days.
Peritoma arborea            Species does not require           2-5
  (Lippitt et al. 1994,       high soil moisture to
  Borders et al. 2008,        germinate.
  Baldwin et al. 2012)
Phacelia ramosissima                                            2
  (Baldwin et al. 2012)
Plantago erecta             Non-dormant, no                     3
  (Gulmon 1992, Montalvo      pre-treatment needed. With
  et al. 2010a, Baldwin       ample water, will
  et al. 2012)                germinate from Sep-Dec
                              with varying temperatures.
Platanus racemosa           Cold moist stratification           2
  (Bonner and Karrfalt        needed to break dormancy.
  2008)
Populus fremontii subs.     Germination is most                 1
  fremontii (Gulmon 1992,     successful at
  Stettler, 1996, Clarke      20-30[degrees]C with
  et al. 2007, Kleinman,      adequate moisture. Seed
  n.d.)                       should not be covered
                              with soil.
Potentilla anserina         Non-dormant. Seeds should be    Short-
  subs. pacifica (Walker      planted in full sun in          lived
  2005, Stevens, n. d.,       lightly packed soil. Keep
  Baldwin et al. 2012)        soil moist.
Pseudognaphalium            Germination stimulated by
  californicum                presence of charred wood
  (Keeley and Keeley          or aqueous extracts of
  1987, Nesom 2013)           charred wood.
Rosa californica            Soak seeds in water                2-4
  (Young 2001i, Lady          overnight prior to sowing.
  Bird Johnson                Seeds germinate slowly,
  Wildflower Center           cold stratification helps
  2007)                       speed process.
Salicornia bigelovii        Irrigate with seawater. Root        1
  (Glenn et al. 1997,         zone salinity (top 15 cm
  Zedler 2001, Baldwin        of soil) should be kept at
  et al. 2012)                a salinity of 70-75 g for
                              high yields.
Salicornia pacifica         Variety S. utahensis grows          2+
  (Khan and Weber 1986,       best at 5% NaCl treatment
  Young 200 lj)               and under temperature
                              regime of 15-5[degrees]C.
Salix exigua subs,          Seeds are non-dormant.             3-4
  exigua (Young and           Optimal germination
  Clements 2003,              temperatures:
  Anderson 2006, Clarke       2-15[degrees]C.
  et al. 2007)
Salix lasiolepis            Sow near soil surface.              3
  (Bonner and Karrfalt        Exposure to light
  2008, Don 2014)             increases germination
                              rate.
Salvia apiana               Scarification and possibly         2-4
  (Stevens 1994,              stratification needed to
  Montalvo and Beyers         break seed dormancy. Sow
  2010d, Baldwin et al.       seed in early fall. Seeds
  2012, Native Plant          may respond to light, so
  Database 2015)              plant in surface soil
                              (1/8-1/4" deep). After
                              planting, soak flats in
                              water.
Salvia mellifera            Physiological dormancy.            1-2
  (Montalvo and Beyers        Exposure to light or
  2010e)                      components of fire
                              (charred wood, smoke,
                              KN03) may stimulate
                              germination.
Sclioenoplectus acutus      Physiological dormancy. Cold        2
  var. occidentalis           stratification breaks
  (Lacroix and Mosher         dormancy. Germination
  1995, Johnson 2004,         rates are low for the
  Baldwin et al. 2012,        species due to the thick
  Baskin and Baskin           pericarp of the achene.
  2014)                       Germination rates increase
                              with overwintering in a
                              pond or water source.
                              Pre-treat seed with 0.05%
                              solution of sodium
                              hypochlorite 5 days prior
                              to sowing. Seeds germinate
                              to a higher percentage
                              when grown in light.
Schoenoplectus              Plant seeds 1/4" under the
  californicus                soil surface. Keep soil
  (Stevens 2003,              surface moist and at a
  Baldwin et al. 2012)        temperature of
                              100[degrees]F.
Spartina foliosa            Best success after cold         4 months
  (Zedler 2001, Baldwin       storage in freshwater.
  et al. 2012)
Stipa cernua                Overheating can kill
  (Laude et al. 1952,         seedlings.
  Amme 2003, Herrera
  et al. 2006)
Stipa lepida                No pretreatment needed.
  (Ashby and Hellmers         Constant light (i.e.
  1959, Elkhom Slough         24-hour photoperiod)
  National Estuarine          exposure is documented to
  Research Reserve 2001,      speed flowering.
  Amme 2003, Dave's
  Garden 2015b)
Suaeda esteroa              Seedlings establish well at         2+
  (Zedler 2001, Baldwin       restoration sites.
  et al. 2012)
Suaeda nigra                Pre-chill recommended.              3
  (Borders, n.d.)
Suaeda taxifolia            Seeds germinate readily with        3
  (Zedler 2001, Baldwin       freshwater irrigation.
  et al. 2012)
Triglochiit maritima        No pre-treatment required.         3-5
  (Young 2002, Baldwin
  et al. 2012, Recon
  Native Plants Inc.
  2015)
Vulpia microstachys         Seeds germinate w/o                4-5
  (Young and Young 1986,      pretreatment. Heating and
  Howard 2006, Baldwin        litter do not increase
  et al. 2012)                germination.

* In instances where published information is insufficient to fill-in
portions of the table, cells are intentionally left blank.

* Seed longevity data from Recon Native Plants Inc. used to
supplement information found in the literature where necessary


Michelle L. Barton (1) *, Ivan D. Medel (2), Karina K. Johnston (2), and Christine R. Whitcraft (1)

(1) California State University Long Beach

(2) Santa Monica Bay Restoration Commission

* Corresponding author: mlbartonl4@gmail.com
Table 1. Suggested field, lab, and greenhouse equipment for seed
collection, cleaning, and germination.

Field equipment                       Lab/greenhouse equipment

Collecting bins/paper bags     Sieves of varying sizes (500 um-2 mm)
Ziploc bags                    Paper envelopes
Pens/pencils/markers           Freezer
Paper clips/binder clips       Refrigerator
Field data collection sheet    Oven
Clipboard                      Growing medium *
Background documentation       Sterile petri dishes *
  (recommended)
Mesh screens/sieves (+)        Flydrogen peroxide
                                 ([H.sub.2][O.sub.2]) *
Tarp(s) (+)                    Nail clippers *
Gloves (+)                     Mothballs *
Gardening shears (+)           Ethylene (ethephon or sliced apple) *
Jepson manual (+)

* = species specific
+ = optional

Table 2. General seed collection method based on plant anatomy
(Wall 2009).

Fruit/seed type                     Collection techniques

Moist fruits/berries   Hand-pluck fruits.
Dehiscent species      Collect entire inflorescences prior to
                         dispersal. Alternatively, secure cloth bags
                         around ripening stalks to capture dispersed
                         seed.
Inflorescences         Strip inflorescences.
Seed heads             Shake ripe seed directly onto a tarp or
                         collection bag underneath the target plant.
Seed clusters          Remove entire seed cluster from plant.

Table 3. Detailed methodology for techniques commonly employed to
break seed dormancy.

Method                              General description

Scarification         Mechanically scar seed coat with sandpaper,
                        knives, files, or clippers. Alternatively
                        soak seed in acid or hot water (Emery 1988,
                        Lippitt 1994, Bonner and Karrfalt 2008).
Hot water treatment   Place seeds into hot water (180-200[degrees]F)
                        and leave them to soak as the water cools
                        (Emery 1988, Bonner and Karrfalt 2008).
Dry heat              Expose seeds to 180-212[degrees]F heat. Use of
                        an incubator, rather than oven, preferred
                        (Emery 1988).
Charate               Expose seeds to ash from burned plants. This
                        may neutralize germination inhibitors in
                        species that naturally germinate when exposed
                        to fire (Emery 1988, Baskin and Baskin 2014).
Fire                  Expose seeds to direct flame. This may be
                        effective as a means to spur germination in
                        species that naturally germinate when exposed
                        to fire (Baskin and Baskin 2014).
Water                 Soak seeds in water to leach out water-soluble
                        inhibitors (Baskin and Baskin 2014).
Cold stratification   Store seeds in cold conditions
                        (35-41[degrees]F) for 1 -3 months to simulate
                        winter conditions (Bonner and Karrfalt 2008,
                        Elsey-Quirk et al. 2009, Baskin and Baskin
                        2014).
Warm stratification   Store seeds in warm conditions (65[degrees]F or
                        higher) (Baskin and Baskin 2014).
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Author:Barton, Michelle L.; Medel, Ivan D.; Johnston, Karina K.; Whitcraft, Christine R.
Publication:Bulletin (Southern California Academy of Sciences)
Article Type:Report
Geographic Code:1USA
Date:Apr 1, 2016
Words:15834
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